Economics 1st Edition Acemoglu

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Chapter 7
Perfect Competition and the Invisible
Hand
Questions
1. All else equal, does a steep or flat demand curve result in higher social surplus? How does steepness
of supply affect social surplus?
Answer: Everything else equal, a steep demand curve will result in higher social surplus. As can be
seen in the following figures, social surplus (shown by the triangles A and B) is higher in the market
with a steep demand curve.

Similarly, other things remaining the same, social surplus (shown by the triangles A and B) is higher
in a market with a steeper supply curve.

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 Chapter 7 | Perfect Competition and the Invisible Hand 70

2. How do economic profits and losses allocate resources in an economy?

Answer: When positive economic profits exist in an industry, resources flow to that industry because
of the profits available. This behavior causes resources to flow from less productive uses to more
productive uses. That is, businesses seek to improve their profits and in so doing, they move
resources into the production of goods and services that society values the highest. Similarly, when
firms face economic losses, they exit the industry and put their resources to more profitable uses.
Again, this allocates resources to industries that produce goods and services that society values the
highest.
3. How will the invisible hand move corn prices in response to:
a. a flood that destroys a great deal of the corn crop?
b. a rise in the price of wheat (a substitute for corn)?
c. a change in consumer tastes away from corn dogs toward hot dogs?
d. an increase in the number of people with access to the corn market?
Answer:
a. Since a great deal of the corn crop has been destroyed by the flood, the supply curve for corn
will shift leftwards and the price of corn will increase.
b. When the price of a substitute such as wheat increases, the demand for corn will increase.
The demand curve will shift rightwards and the price of corn will increase.
c. Corn (not surprisingly) is an important input in the production of corn dogs. If consumers
prefer fewer corn dogs, the demand curve for corn will shift leftwards and the price of corn
will fall.
d. As more buyers enter the market, the demand for corn will increase. The demand curve will
shift rightwards and the price of corn will increase.
4. Hardware stores charge higher prices for snow shovels after a big snowstorm. What role do prices
play in the snow shovel market?
Answer: Prices allocate scarce resources (such as snow shovels after a big snow storm) to the
consumers who value snow shovels the highest. Thus prices play an important role in the
maximization of social surplus. Prices in this case will also increase producer surplus (i.e, the
hardware stores are better off as a result of the higher prices) but may decrease consumer surplus (i.e.,
consumers are worse off as a result of the higher prices). The higher prices also provide an incentive
for hardware stores to try to acquire more shovels to sell and to hold a large inventory of snow
shovels at the start of the winter; these decisions will increase social surplus.
5. The market for economics textbooks is in equilibrium. The government decides to relax export
restrictions on paper, leading to an increase in the demand for paper. How does social surplus in the
market for textbooks change? Why? Present a diagram as part of your explanation.
Answer: An increase in the demand for paper would also lead to an increase in the price of paper. As
the price of paper increases, the cost of producing a textbook will increase. This will shift the supply
curve for textbooks to the left. The equilibrium price of a textbook will increase while the equilibrium
quantity will fall. Consumer surplus and social surplus must fall; it is not clear if producer surplus
will rise or fall.

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71 Acemo
oglu, Laibson, and List | Micrroeconomics

The diagram ab
T bove shows demand
d and supply
s in the textbook
t markket. The increease in the price of
paaper shifts thee supply curvve for textbooks from S1 to S2. At the oldd price of papper consumer
suurplus was A + B + C + E,, producer surrplus was F + G + H, and social
s surpluss was A + B + C +
E + F + G + H. H At the new higher
h f paper conssumer surpluss is A, produccer surplus is B +
price for
F, and social su urplus is A + B + F. Thereefore consumeer surplus andd social surpluus both fell; the
t
im
mpact on prod ducer surplus is ambiguouss.
6. What could explain n why South Korea’s gross domestic prroduct (GDP) per capita inncreased so muuch
fasterr since the 197
70s than Nortth Korea’s GD
DP per capitaa?
Answeer: One of thee factors that could explainn the divergennce between SouthS Korea’s and North
Koreaa’s per capita GDP is the factfa that Southh Korea is a market
m econommy while Norrth Korea is a
comm mand economy y. The price mechanism
m alllocates resouurces in markeet economies to their mostt
efficieent use. It also aligns the inncentives of sellers
s and buuyers. Centrall planners in command
c
econoomies, howev ver, face coorddination and incentive
i probblems. The ceentral plannerr does not fullly
underrstand consum mer wants andd the productiion capabilitiees of every seector of the ecconomy, and it i is
difficuult to providee workers withh appropriatee incentives.
7. In a command econ nomy, a plannning agency sets
s prices forr various inpuuts and final goods.
g In a market
econoomy, supply and
a demand decide
d the pricces of variouss goods. In booth cases, therre is a set of prices
p
operaating in the economy. Thenn why are marrket economiees consideredd more efficient than plannned
econoomies?
Answeer: Although planning ageencies set pricces, they cannnot always acccurately alignn the interests of
buyerrs and sellers or solve the coordination
c p
problem of brringing agentss together to trade.
t Any
indiviidual can onlyy know a smaall fraction off all that is knoown collectivvely and so a planning
p agenncy
cannoot replicate the work of thee invisible hannd. In a markeet economy on o the other haand, the pricee
mechaanism ensures that econom mic agents maake trades thaat are in their best
b interest anda maximizee
sociall surplus.

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 Chapter 7 | Perfect Competition and the Invisible Hand 72

8. If your professor decided to give all of his students the highest grade in the class, would that affect
your classmates’ incentives to study?
Answer: If every student were given the same grade irrespective of how he or she did on an exam,
students’ incentive to study. Even without studying, a student would be ‘guaranteed’ a higher grade
than he would have received otherwise. Eventually, one can expect students in the class to stop
studying altogether.
9. Sofia, a political science student, thinks that the government should intervene to revive declining
industries like video rentals and print newspapers. The government, she reasons, can resolve the
coordination problem of getting the agents in these markets to trade. Do you agree with her? Explain
your answer.
Answer: Sofia’s reasoning is flawed because declining industries do not face a coordination problem.
A coordination problem results when economic agents with coinciding interests cannot be brought
together to trade. However, in this case, falling demand is responsible for the decline of industries like
video rentals and print newspapers. Even if the government attempted to bring together economic
agents in these industries, it is unlikely that demand for rented videos and newspapers will increase.
10. Are all efficient outcomes also equitable? Explain.
Answer: Equity is concerned with the distribution of resources across society. An efficient outcome is
one that maximizes social surplus. However, maximizing social surplus is not always consistent with
equity considerations. An economy can make the most efficient use of the resources that it has but at
the same time, these resources may not be equally distributed in society.
11. Are there real-world markets that resemble double oral auctions? Suppose you had to organize a
double oral auction for a good that has perfectly elastic demand. Do you expect prices to approach the
competitive equilibrium?
Answer: Double oral auctions are similar to how trading actually works on stock exchanges—traders
announce bids and asks and if they match, a trade is executed.
Double oral auction experiments with many different market variants - including varying the
elasticity of supply and demand and the numbers of buyers and sellers - have shown that the
equilibrium price in the market will be very close to where the supply and demand curves intersect.
So, the price in a double oral auction for a good with perfectly elastic demand is also expected to
approach the equilibrium price.
12. Imagine you are a buyer in a double oral auction with a reservation value of $10 and there is a seller
asking $8.
a. How much will you gain from accepting this offer?
b. If you are the only buyer, and you know that the lowest ask price is $2, should you accept this
offer?
Answer:
a. By accepting this offer, you will gain $2 ($10 – $8).
b. Yes. By accepting this offer, you will gain $8 ($10 – $2). If you choose to accept an offer
from a different seller, your surplus will be lower. By accepting the lowest ask price of $2,
you will maximize your surplus from this trade.

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73 Acemo
oglu, Laibson, and List | Micrroeconomics

Problem
ms
1. The following
fo diaggram shows thhe market dem mand and maarket supply for fo sweaters. Calculate
C
consuumer surplus, producer surpplus, and sociial surplus in this market.

Answeer: Producer surplus is thee area of trianggle I and is eqqual to ½ x 1000 x (60 – 20) = 2000.
Consuumer surplus is the area off triangle II annd is equal to ½ x 100 x (90 - 60) = 15000. Social surpplus
is the sum of consu
umer surplus and producerr surplus and is equal to 15500 + 2000 = 3500.

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 Chapter 7 | Perfect Competition and the Invisible Hand 74

2. Look at Exhibit 7.1 in the chapter that shows the reservation values of the buyers and sellers in the
iPod market. Suppose trades are arranged in this market such that everyone can make a trade without
losing money. So, Madeline buys from Fiona at a price of $70, Katie buys from Matt at a price of
$60, Sean buys from Adam at a price of $50 and so on.
Reservation Values of Buyers and Sellers in the iPod Market
Buyers Value($) Sellers Cost ($)
Madeline 70 Tom 10
Katie 60 Mary 20
Sean 50 Jeff 30
Dave 40 Phil 40
Ian 30 Adam 50
Kim 20 Matt 60
Ty 10 Fiona 70
Since everyone who wants an iPod obtains one, and everyone who wants to get rid of their iPod sells
it at the price they wanted, is social surplus maximized in the market?
Answer: When trades are arranged in the market such that the buyer’s reservation values are matched
with the minimum price that the sellers are willing to accept, the social surplus in the market is equal
to zero. For example, Madeline buys an iPod from Fiona for $70. Since this is the maximum that
Madeline was willing to pay and the minimum that Fiona was willing to accept, there is no producer
surplus or consumer surplus from this trade. This will hold for all the trades between the other
participants and the social surplus will be zero.
Buyers Value($) Sellers Cost ($)
Madeline 70 Fiona 70
Katie 60 Matt 60
Sean 50 Adam 50
Dave 40 Phil 40
Ian 30 Jeff 30
Kim 20 Mary 20
Ty 10 Tom 10
3. There are four consumers willing to pay the following amounts for an electric car:
Consumer 1: $70,000 Consumer 2: $20,000 Consumer 3: $80,000 Consumer 4: $40,000
There are four firms that can produce electric cars. Each can produce one car at the following costs:
Firm A: $30,000 Firm B: $60,000 Firm C: $40,000 Firm D: $20,000
Each firm can produce at most one car.
Suppose we wanted to maximize the difference between consumers’ willingness to pay for electric
cars and the cost of producing those cars, i.e., we wanted to maximize social surplus.
a. How many electric cars should we produce?
b. Which firms should produce those cars?

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75 Acemoglu, Laibson, and List | Microeconomics

c. Which consumers should purchase those cars?

d. Find the maximum social surplus in the electric car market.
Answer:
a. We should produce three cars. The value of the fourth car would be just $20,000 and would
cost $60,000 to produce.
Consumers Firms
(I) (II)
$80,000 3 $20,000 D
$70,000 1 $30,000 A
$40,000 4 $40,000 C
$20,000 2 $60,000 B
b. The three lowest cost firms should produce: Firms A, C, and D
c. The three consumers who value electric cars the highest should own electric cars: Consumers
1, 3, and 4.
d. Willingness to pay of the consumers = $80,000 + $70,000 + $40,000 = $190,000
Total Cost = $20,000 + $30,000 + $40,000 = $90,000
Social surplus = $190,000 – $90,000 = $100,000
4. Let us continue with the electric car example from problem 3. Suppose the market for electric cars is
competitive.
a. Show that the equilibrium price in this market is $40,000.
b. Which firms will produce an electric car if the price is $40,000?
c. Which consumers will buy an electric car when the price is $40,000?
d. Calculate consumer, producer surplus, and social surplus when the price is $40,000.
e. Compare your answers to those for problem 3.
Answer:
a. When the price is $40,000, three consumers (all except 2) are willing to buy an electric car
because their willingness to pay is greater than or equal to price. Three firms (all except B)
are willing to sell an electric car since their cost is less than or equal to the price. Therefore
supply equals demand when the price is $40,000.
b. Firms A,C, and D
c. Consumers 1,3 and 4
d. Consumers 1, 3, and 4 are willing to pay a total of $80,000 + $70,000 + $40,000 = $190,000
for three cars. At a price of $40,000 they will pay 3 x $40,000 = $120,000 for those cars.
Therefore consumer surplus equals $190,000 - $120,000 = $70,000. It will cost Firms D, A,
and C a total of $20,000 + $30,000 + $40,000 = $90,000 to produce those cars. At a price of
$40,000 their revenues equal 3 x $40,000 = $120,000 Therefore producer surplus equals
$120,000 - $90,000 = $30,000. Social surplus is the sum of consumer and producer surplus.
Therefore social surplus equals $70,000 + $30,000 = $100,000.

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 Chapter 7 | Perfect Comp
petition and thee Invisible Han
nd 76

e.. The point here

h is that coompetitive maarkets lead too efficiency. The
T market ouutcome here leads
to the max
ximum social surplus we foound in probleem 3.
5. The following
fo figu
ure shows the demand and supply of teleevision sets inn a city. Sincee TVs are
considdered normal goods, demaand increases from D1 to D2 in response to an increasse in the incom
me
level.

a. Use
U the figure to complete the
t table beloow:
Befo
ore Income Ro
ose After Income
I Rose Chaange

Co
onsumer Surp
plus

Prroducer Surplus

So
ocial Surplus

b. Use
U your answ
wers to part (a) of this problem to answeer the followinng questions:
i. Did coonsumer surpllus definitely rise, definitely remain connstant, definittely fall, or is the
directiion of the chaange in consum
mer surplus unclear?
u
ii. Did prroducer surpluus definitely rise,
r definitelyy remain constant, definiteely fall, or is the
t
directiion of the chaange in produccer surplus unnclear?
iii. Did so
ocial surplus definitely
d risee, definitely reemain constannt, definitely fall, or is the
directiion of the chaange in social surplus uncleear?

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77 Acemoglu, Laibson, and List | Microeconomics

Answer:
a. The completed table is as follows:
Before Income Rose After Income Rose Change

Consumer Surplus A+C A+B B–C

Producer Surplus F F+C+D C+D

Social Surplus A+C+F A+B+F+C+D B+D

b. As shown in the table:

i. The change in consumer surplus is B – C. We do not know if B is greater than or less
than C and so we do not know if consumer surplus rose or fell.
ii. Producer surplus rises by C + D.
iii. Social surplus rises by B + D.
6. The market for electric drills in a certain country is characterized by a large number of buyers and
sellers and every buyer who wants a drill and can afford one has bought one. In other words, the
market for drills is in equilibrium.
a. Does this also mean that it is Pareto efficient? Explain your answer.
b. If some of the buyers in this market are now willing to pay more than they did earlier, would your
answer change?
c. Compared to the market for cars, the market for vintage buttons has fewer buyers and sellers.
Social surplus is likely to be higher in the market for cars than in the vintage button market. Is it
then correct to assume that the outcome in the car market is Pareto efficient while in the vintage
button market it is not? Explain.
Answer:
a. An outcome is Pareto efficient if no individual can be made better off without making
someone else worse off. A competitive market maximizes social surplus and so the market
outcome is Pareto efficient. Assuming that none of the large number of sellers has enough
market share to influence price, the market for electric drills is competitive, social surplus is
at its maximum and so, the market outcome is Pareto efficient.
b. The level of social surplus might increase if buyers are now willing to pay more than they did
earlier but as long as the competitive market equilibrium holds, the market outcome is still
Pareto efficient.
c. While social surplus in the market for vintage buttons is lower than in the market for cars, it
is incorrect to assume that the equilibrium in the car market is Pareto efficient while the
equilibrium in the vintage button market is not. The size of the social surplus in the market
does not determine whether the market is Pareto efficient or not. Pareto efficiency would only
depend on whether an individual can be made better off without making someone else worse
off.

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 Chapter 7 | Perfect Competition and the Invisible Hand 78

7. The following tables show a small firm’s long-run average cost of manufacturing a good at two
different plants:
Plant 1
Quantity Total Cost Average Cost Marginal Cost
1 50
2 106
3 164
4 224
5 287
6 355
7 430
8 520
9 618

Plant 2
Quantity Total Cost Average Cost Marginal Cost
1 20
2 52
3 90
4 130
5 175
6 227
7 285
8 345
9 407

a. Complete the third and fourth columns of each table.

b. Suppose the price of the good is $60. How much should the firm produce in each plant in order to
maximize the firm’s profit? Find the firm’s profit.
c. A new manager is assigned to the production department. He thinks that the firm can profitably
move all production to Plant 2 since the average cost of production is lower in Plant 2 than in
Plant 1. If the firm only uses Plant 2, how much should it produce in order to maximize profits?
Find the firm’s profit. Assume zero fixed cost.

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79 Acemoglu, Laibson, and List | Microeconomics

Answer:
a.
Plant 1
Quantity Total Cost Average Cost Marginal Cost
1 50 50.00 50
2 106 53.00 56
3 164 54.67 58
4 224 56.00 60
5 287 57.40 63
6 355 59.17 68
7 430 61.43 75
8 520 65.00 90
9 618 68.67 98

Plant 2
Quantity Total Cost Average Cost Marginal Cost
1 20 20.00 20
2 52 26.00 32
3 90 30.00 38
4 130 32.50 40
5 175 35.00 45
6 227 37.83 52
7 285 40.71 58
8 345 43.13 60
9 407 45.22 62
b. Since the price of the good is $60, profits are maximized when marginal cost of production is
equal to $60 at both plants. This occurs when 4 units are produced at Plant 1 and 8 units are
produced at Plant 2. The firm’s revenue will be 12 x $60 = $720, its total costs will be $224 +
$345 = $569, and it will earn a profit of $720 - $569 = $151.
c. If the firm uses just Plant 2 it would maximize profits by producing just 8 units (since price
equals marginal cost in Plant 2 at 8 units). Its revenue will be 8 x $60 = $480, its cost will be
$345, and its profit will be $480 - $345 = $135.

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 Chapter 7 | Perfect Comp
petition and thee Invisible Han
nd 80

8. The following
fo figu
ure shows a firm’s marginaal and averagee costs of production.

a. T
The equilibrium
m price in thiis market is $5. At this pricce, does the fiirm earn proffits or is it makking
loosses?
b. Frrom the givenn informationn, can you connclude whetheer the firm is operating in a competitivee
m
market? Explaain your answer.
c. The
T price of th
he good increaases to $8. Hoow does this change
c your answer
a to parrts a and b?
Answeer:
a.. At a price of $5, the firm
m produces 60,000
6 units of
o the good. Itt does not earnn profits or loosses.
nue from 60,0000 units = $3300,000
Total reven
Total costss of producingg 60,000 unitts = $300,000
Since totall costs = total revenues, proofits are equaal to zero.
b.. Firms in co
ompetitive markets
m earn zeero economicc profits in thee long run. Att a price of $55, this
firm earns zero econom
mic profits. Soo from the givven informatioon, we can coonclude that thhe
firm may be
b operating in
i a competitiive market.
c.. If the mark
ket price increeases to $8, thhe firm will produce
p 80,0000 units of thee good.
Total reven
nue from 80,0000 units = $6640,000
Total costss of producingg 80,000 unitts = $440,000
Since total revenuee exceeds totaal costs by $2200,000, the firm
fi earns ecoonomic profitss. In the long run,
competitive firms earn
e zero ecoonomic profitss. This firm makes
m positivee economic prrofits when thhe
price is $8. This is however, nott sufficient too conclude thaat the firm is not
n operating in a competitive
markeet. This firm might
m be operrating in the short
s run wheere new firms have not enteered the markket
and coompeted awaay its profits.

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81 Acemo
oglu, Laibson, and List | Micrroeconomics

9. Hospiitals in City A are profit maximizing

m peerfectly competitive firms. Hospitals in City B, on the
other hand, are run n by non-profifit charities that try to minim
mize the longg-run average cost of treatiing
patiennts. Hospitals in both citiess have the sam me average annd marginal cost.
c Show thaat hospitals inn both
cities will be the saame size.
Answeer: Hospitals in City B treaat the numberr of patients thhat minimizees long run avverage cost
(LRAAC). Hospitalss in City A wiill equate pricce and marginnal cost in ordder to maximiize profits andd
thereffore, the equillibrium price must equal loong run marginal cost (LRRMC). Entry will
w eliminate
(econnomic) profits and so in equuilibrium, price must also equal LRAC.. When LRAC C equals LRMMC,
LRAC C is minimizeed. So, for hospitals in Cityy A, price is equal
e to LRM
MC which is eqqual to the
minimmum point of LRAC. Therefore, hospitaals in both cities will buildd hospitals thaat treat the samme
numbber of patientss.

In thee diagram abo

ove, hospitals in both citiess will serve Q1 patients.

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 Chapter 7 | Perfect Comp
petition and thee Invisible Han
nd 82

10. The equilibrium

e reent in a town is
i $500 per month
m and the equilibrium number
n of apartments is 1000.
The city
c now passees a rent contrrol law that seets the maximmum rent at $4400. The diaggram below
summ
marizes the sup pply and demmand for aparttments in thiss city.

a. Use
U the figure to complete the
t table beloow

Befo
ore Rent Conttrol After Rent
R Control Chaange
Co
onsumer Surp
plus

Prroducer Surpllus

So
ocial Surplus

b. Use
U your answ
wers to part (a) of this problem to answeer the followinng questions:
i. Did coonsumer surpllus definitely rise, definitely remain connstant, definittely fall, or is the
directiion of the chaange in consum
mer surplus unclear?
u
ii. Did prroducer surpluus definitely rise,
r definitelyy remain constant, definiteely fall, or is the
t
directiion of the chaange in produccer surplus unnclear?
iii. Did so
ocial surplus definitely
d risee, definitely reemain constannt, definitely fall, or is the
directiion of the chaange in social surplus uncleear?

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83 Acemoglu, Laibson, and List | Microeconomics

Answer:
a. The completed table is as follows:

Before Rent Control After Rent Control Change

Consumer Surplus I + II I + III III – II

Producer Surplus III + IV + V V – III – IV

Social Surplus I + II + III + IV + V I + III + V – II – IV

b. As shown in the table:

i. The change in consumer surplus is III – II and so it is not clear if consumer surplus rose
or fell. The consumers who rent those 60 apartments that are available under rent control
are better off because they now pay $400 per month instead of $500; this benefit is area
III. But the people who cannot find an apartment under rent control are worse off. They
lose the consumer surplus they realized from the 40 apartments that are no longer
available as a result of rent control; this loss is area II.
ii. Suppliers are unambiguously worse off as a result of price controls. They receive $400
per month instead of $500 per month for each of the 60 apartments they continue to offer
under rent control (area III) and they lose the producer surplus they realized from the 40
apartments they no longer offer (area IV).
iii. Total surplus falls by the sum of II (lost consumer surplus from the 40 apartments that are
no longer offered) and IV (lost producer surplus from those 40 apartments).
11. According to reports in the Chinese media, commuters in Beijing are facing a somewhat paradoxical
situation: they find it difficult to get a cab while hundreds of cabs lie idle during rush hour. The
demand for taxis in Beijing has increased as average incomes have risen. Government-determined
gasoline prices have also increased. But the government, worried about rising prices, has left the fares
cabs can charge their customers unchanged.
a. Use supply and demand curves to explain what has happened in the market for cabs in Beijing.
b. Based on your understanding of how the invisible hand works, what do you think should be done
to correct this problem?
Answer:
a. The following diagram shows the demand and supply curves for cabs in Beijing. The increase
in demand causes the demand curve to shift to the right. Since gasoline prices have increased,
the cost of running a cab in Beijing has also increased. This will shift the supply curve to the
left. The equilibrium quantity may or may not change depending on the degree to which both
curves shift but the equilibrium price in the market will increase. However, since the
government does not allow cab fares to increase, the number of cabs demanded exceeds the
supply. This explains why commuters are unable to find a cab when they need one.

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 Chapter 7 | Perfect Comp
petition and thee Invisible Han
nd 84

As can be seen in the figure, the incrrease in demaand and decrease in supply have actuallyy
increased the
t equilibriuum price from m P to P’. The equilibrium quantity
q shouuld also have
increased from
f Q to Q’. The distancee between Q0 and Q1 show ws the extent by
b which dem mand
exceeds su
upply as a resuult of the pricce control.

Market for Cab Rides

R

b.. An increasse in demand combined wiith a decreasee in supply im mplies an increease in price.
Therefore, to fix this prroblem, the caab fare must be
b allowed to increase. Forr the market too be
in equilibrrium, fares muust be allowedd to increase to P’. At this price, quantitty supplied will
w
equal demand at the quaantity demandded.
See htttp://www.economist.com//node/215515537 and
http:///www.theatlaantic.com/inteernational/archhive/2013/01/its-harder-thhan-ever-to-caatch-a-cab-in--
beijinng/267239/
12. The following
fo quote is from a section on foood shortages inn a book on thhe Soviet ecoonomy:
"Why there is no fiish ... I can't imagine,"
i wroote one indignnant citizen too Anastas Mikkoyan, head of
o the
Food Ministry, in 1940.
1 "We haave seas, and they are still the same as before,
b but theen you could have
y wanted off whatever kinnd, and now I have even foorgotten whatt it looks like."
as muuch [fish] as you
Industries and agriiculture in thee former Soviet Union were state-controolled and the economy’s
e
resourrces were allo
ocated by a ceentral agencyy, Gosplan. Inn the above paassage, the citizen cannot
underrstand why there is a shortaage of fish altthough the coountry possessses the same resources
r thatt it
did beefore the econ
nomy transitiooned to centraal planning. What
W could exxplain this ouutcome?
Answeer: The shortaage of fish coould be explaiined by the faact that the economy was centrally
c plannned.
A cenntral planningg agency is unnlikely to corrrectly estimatee the demandd for various goods,
g in this case,
fish. It
I is also not likely
l to be abble to match production
p caapabilities witth the econom
my’s resourcess. In
other words, since central plannning agencies cannot repliccate the work of the invisibble hand,
mismmatches betweeen demand annd supply are likely to exisst in the econoomy.
From: Everyday Sttalinism: Orddinary Life in Extraordinar
E ry Times: Sovviet Russia in the 1930s (Shheila
Fitzpaatrick)

© 2015 Peearson Education

n, Inc.
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Monogonopora, in which a single orifice serves for both male and
female organs, may be taken the common garden snail (Helix
aspersa), the accompanying figure of which is drawn from two
specimens found in the act of union (Fig. 53).
Fig. 53.—Genitalia of Helix aspersa Müller,
drawn from two individuals in the act of
union, from a dissection by F. B. Stead.
 A.G, albumen gland.
C, coecum.
Cr, crop.
D.S, dart sac.
E, eye (retracted).
Fl, flagellum.
H.D, hermaphrodite duct.
H.DF, ditto, female portion.
H.DM, ditto, male portion.
H.G, hermaphrodite gland.
L, liver.
M.G, M.G, mucous glands.
Ov, oviduct.
P.S, penis sac.
R.M, retractor muscle of penis.
Sp, spermatheca.
V, vagina.
V.D, vas deferens.

Beginning from the inside and proceeding outwards we have

firstly the hermaphrodite gland or ovo-testis (h.g.), a yellowish white
mass of irregular shape, embedded in the liver (l.) and forming part
of its spiral but not reaching quite to the apex. Within this gland are
developed the ova and spermatozoa. The former are rather large
round cells, produced within the outer wall of the gland, while the
spermatozoa, which are produced in the more central part, are
thread-like bodies, generally aggregated in small bundles. From the
hermaphrodite gland the ova and spermatozoa pass through the
upper part of the hermaphrodite duct (h.d.), which is always more or
less convoluted. Below the convoluted portion, the duct opens into
the albumen gland (a.g.), a large linguiform mass of tissue which
becomes dilated at the time of pairing, and secretes a thick viscid
fluid which probably serves to envelop the ova. Up to this point both
the male and female elements follow the same course, but on their
exit from the albumen gland they diverge. The hermaphrodite duct
becomes greatly enlarged, and is partially divided by a kind of
septum into a male and female portion. These run parallel to one
another, the larger or female portion (h.df.), through which the ova
pass (and which is sometimes termed the uterus) being dilated into a
number of puckered folds, while the smaller or male portion (h.dm.)
is comparatively narrow, and not dilated. At their anterior end, the
two portions of the duct separate completely from one another, the
female portion being then termed the oviduct (ov.) and the male
portion the vas deferens (v.d.).
Following first the oviduct, we find that it soon widens into the
vagina (v.), which is furnished with a pair of mucous glands (m.g.),
one on each side. These are much branched, and resemble little
bunches of whitish seaweed. A little above the mucous glands a long
tube diverges from the vagina, which is furnished with a produced
coecum (c.) and a pouch, the spermatheca (sp.) at the extreme end.
In this pouch, and in the duct leading to it, is stored the
spermatophore received in union with another snail. Just below the
mucous glands the vagina is joined by the dart sac (d.s.), which is
more fully described below. Finally, at its lower end the vagina unites
with the penis sac at a point just posterior to the common orifice.
Returning now to the male organs, we find that the vas deferens
is the continuation of the male portion of the hermaphrodite duct,
after its final separation from the female portion. It passes under the
retractor muscle of the upper right tentacle, which has been cut away
in the specimen figured, to dissect it out. Just before the vas
deferens widens into the penis sac, it branches off into a long and
tapering tube, the flagellum, in which the spermatozoa are stored
and become massed together in the long packet known as the
spermatophore. The penis sac (p.s.) is the continuation of the vas
deferens beyond the point at which the flagellum diverges. It joins
the vagina at its extreme anterior end, uniting with it to form the
common genital aperture, which cannot be exactly represented in
the figure. The penis itself lies in the interior of the penis sac, and is
a rather long muscular tube which is protruded during union, but at
other times remains retracted within the sac.
In the Helicidae generally, the form of the generative organs
varies with each separate species, sometimes merely as regards the
size of the different parts, at others in the direction of greater
simplicity or complication. The mucous glands may be absent, and
the flagellum greatly reduced in size, or absent altogether.
 The Dart Sac.—A remarkable part of the reproductive system in
many of the true Helicidae is the so-called dart, Liebespfeil, or telum
veneris. It consists of ‘a straight, or curved, sometimes slightly
twisted tubular shaft of carbonate of lime, tapering to a fine point
above, and enlarging gradually, more often somewhat abruptly, to
the base.’ The sides of the shaft are sometimes furnished with two or
more blades; these are apparently not for cutting purposes, but
simply to brace the stem. The dart is contained in a dart sac, which
is attached as a sort of pocket to the vagina, at no great distance
from its orifice. There are four different forms of sac. It may be single
or double, and each of these divisions may be bilobed, each lobe
containing one dart at a time. In Helix aspersa the dart is about 5/16
in. in length, and ⅛ in. in breadth at its base (see Fig. 54).
It appears most probable that the dart is employed as an adjunct
to the sexual act. Besides the fact of the position of the dart sac
anatomically, we find that the darts are extruded and become
embedded in the flesh just before or during the act of copulation. It
may be regarded, then, as an organ whose punctures induce
excitement preparatory to sexual union. It only occurs in well-grown
specimens. When once it begins to form, it grows very rapidly,
perhaps not more than a week being required for its entire formation.

Fig. 54.—Darts of British land snails: A,

Hyalinia excavata Bean; B, Helix
hortensis Müll.; C, Helix aspersa
Müll. (After Ashford.)
 The dart is almost confined to Helicidae, a certain number of
exceptions being known which border on Helix. Hyalinia nitida and
excavata are the only British species, not Helices, which are known
to possess it. It has not been noticed to occur in the slugs, except in
the N. American genus Tebennophorus. About one-third of the
British Helices are destitute of the dart.[260] H. rufescens possesses
a double bilobed sac, but only two darts, which lie in the lower lobes.
It does not use the darts, and could not do so, from the relative sizes
of dart and sac; it has often been watched when uniting, but the use
of the darts has never been observed. From this it has been inferred
that the darts are degenerate weapons of defence, and that they
were in fact at one time much stronger organs and more often used.
[261] This theory, however, does not seem consistent with the whole
circumstances of the occurrence, position, and present use of the
darts.
Hermaphrodite Mollusca.—(b) Digonopora.—As an example of
the Digonopora or hermaphrodite Mollusca with separate generative
apertures for the male and female organs, we may take the common
Limnaea stagnalis (Fig. 55). It will be seen from the figure that the
relative positions of the hermaphrodite gland and duct, and of the
albumen gland, are the same as in Helix. When the oviduct parts
company from the vas deferens, it becomes furnished with several
accessory glands, one of which (Gl.E.) probably serves as a
reservoir for the ova, and answers more or less to a uterus. The tube
leading to the spermatheca is short, and there is no divergent
caecum. The female orifice lies near to the external opening of the
branchial cavity. The vas deferens, which is very long, is furnished
with a large prostate gland. The penis sac is greatly dilated, and
there is no flagellum. The male orifice is behind the right tentacle,
slightly in advance of the female orifice (compare Fig. 102).
Fig. 55.—Genitalia of Limnaea stagnalis
L. (from a dissection by F. B. Stead),
× 2.
 A.G, albumen gland.
Ac.G, accessory gland.
F.O, female orifice.
Gl.E, glandular enlargement.
H.D, hermaphrodite duct.
H.G, hermaphrodite gland.
Li, liver.
M.O, male orifice.
P, penis sac.
Pr, prostate.
R.M, retractor muscle of penis.
Sp, spermatheca.
V.D, vas deferens.]

Most of the Opisthobranchiata, but not all, have separate sexual

orifices. Numerous variations from the type just described will be
found to occur, particularly in the direction of the development of
accessory glands, which are sometimes very large, and whose
precise purpose has in many cases not been satisfactorily
determined.
Pelecypoda.—In the dioecious Pelecypoda, which form the great
majority, the reproductive system is simple, and closely parallel in
both sexes. It consists of a pair of gonads, which are either ovaries
or testes, and a pair of oviducts or sperm-ducts which lead to a
genital aperture. The gonads are usually placed symmetrically at the
sides or base of the visceral mass. The oviduct is short, and the
genital aperture is usually within the branchial chamber, thus
securing the fertilisation of the ova by the spermatozoa, which are
carried into the branchial chamber with the water which passes
through the afferent siphon.
Hermaphrodite Pelecypoda are rare, the sexes being usually
separate. The following are assured instances: Pecten glaber, P.
jacobaeus, P. maximus, Ostrea edulis, Cardium norvegicum,
Pisidium pusillum, Cyclas cornea, Pandora rostrata, Aspergillum
dichotomum, and perhaps Clavagella. The greater number of these
have only a single genital gland (gonad) on each side, with a single
efferent duct from each, but part of the gland is male and part
female, e.g. in the Pectens above mentioned. Pandora and
Aspergillum have two distinct glands, respectively male and female,
on each side, each of the two glands possessing its separate duct,
and the two ducts from each side eventually opening near one
another. It appears probable that the Septibranchiata (Cuspidaria,
Poromya, Lyonsiella, etc.) must also be added to the number of
hermaphrodite Pelecypoda which have separate male and female
glands.
It is worthy of remark that all the hermaphrodite Pelecypoda
belong to forms decidedly specialised, while forms distinctly
primitive, such as Nucula, Solenomya, Arca, and Trigonia are all
dioecious. In Gasteropoda similarly, the least specialised forms (the
Amphineura, with the exception of the Neomeniidae, and the
Rhipidoglossa) are dioecious. It is possible therefore that in the
ancestors of the Mollusca the separation of the sexes had already
become the normal type of things, and that hermaphroditism in the
group is, to a certain extent, a sign or accompaniment of
specialisation.[262]
Development of Fresh-water Bivalves.—The vast majority of
fresh-water bivalves either pass the larval stage entirely within the
mother, and do not quit her except in a perfectly developed form
(Cyclas, Pisidium), or assume a mode of development in which free
larvae indeed occur, but are specially modified for adaptation to
special circumstances (Unio). Cyclas and Pisidium, and no doubt all
the kindred genera, preserve their ova in a sort of brood-pouch
within the gills, in which the ova pass the earlier stages of their
development. But, even so, the larva of these genera retains some
traces of its original free-swimming habits, for a rudimentary velum,
which is quite useless for its present form of development, has been
detected in Cyclas.
The larva of Dreissensia (see Fig. 47, A), so far as is at present
known, stands alone among fresh-water bivalves in being free-
swimming, and to this property has been attributed, no doubt with
perfect justice, the fact of the extraordinarily rapid spread of
Dreissensia over the continent of Europe (chap. xvi.). In expelling the
ova, the parent slightly opens the shells and then quickly closes
them, shooting out a small point of white slime, which is in fact a little
ball of eggs. The general course of development is precisely parallel
to that of marine Pelecypoda, greatly resembling, so far as form is
concerned, certain stages in the growth of the larvae of Modiolaria
and Cardium, as figured by Lovén.[263]
In June and July the larvae appear in large numbers on the
surface of the water, when in spite of their exceedingly small size,
they can be captured with a fine hand-net. They pass about eight
days on the surface, feeding apparently on minute floating algae.
During this time, the principal change they undergo is in the
formation of the foot, which first appears as a small prominence
midway between the mouth and anus, and gradually increases in
length and flexibility. When the larva sinks to the bottom, the velum
soon disappears entirely, the foot becomes exceedingly long and
narrow, while the shell is circular, strongly resembling a very young
Cyclas.
Larvae of Unionidae.—The early stages of the development of
Unio and Anodonta (so far as the species of North America, Europe,
and Asia are concerned) is of extreme interest, from the remarkable
fact that the young live for some time parasitically attached to certain
species of fresh-water fishes. In order to secure this attachment, the
larva, which is generally known as Glochidium, develops a long
filament which perhaps renders it aware of the neighbourhood of a
fish, and also a larval shell furnished with strong hooks by which it
fastens itself to the body of its unconscious host (Fig. 56). According
to some interesting observations made by Mr. O. H. Latter,[264] the
ova pass into the external gill of the mother, in which is secreted a
nutritive mucus on which they are sustained until they arrive at
maturity and a suitable opportunity occurs for their ‘being born.’ If
this opportunity is deferred, and the Glochidia mature, their so-called
‘byssus’ becomes developed, and by being entangled in the gill
filaments of the parent, prevents their escaping. It is interesting to
notice that, when the nutritive mucus of the parent is used up, it
becomes, as it were, the turn of the children to provide for
themselves a secondary mode of attachment.
 Fig. 56.—A, Glochidium immediately after it is hatched: ad,
adductor muscle; by, ‘byssus’ cord; s, sense organs; sh,
shell. B, Glochidium after it has been on the fish for some
weeks: a.ad, p.ad, anterior and posterior adductors; al,
alimentary canal; au.v, auditory vesicle; br, branchiae; f,
foot; mt, mantle. (Balfour.)
The mother Anodonta does not always retain the Glochidium until
fish are in her neighbourhood. Gentle stirring of the water caused
them to emit Glochidium in large masses, if the movement was not
so violent as to cause alarm. The long slimy masses of Glochidium
were observed to be drawn back again within the shell of the mother,
even after they had been ejected to a distance of 2 or 3 inches.
It is a mistake to assert that the young Glochidium can swim.
When they finally quit the mother, they sink to the bottom, and there
remain resting on their dorsal side, with the valves gaping upwards
and the so-called byssus streaming up into the water above them.
There they remain, until a convenient ‘host’ comes within reach, and
if no ‘host’ comes within a certain time, they perish. They are
evidently peculiarly sensitive to the presence of fish, but whether
they perceive them by smell or some other sense is unknown. “The
tail of a recently killed stickleback thrust into a watch-glass
containing Glochidium throws them all into the wildest agitation for a
few seconds; the valves are violently closed and again opened with
astonishing rapidity for 15–25 seconds, and then the animals appear
exhausted and lie placid with widely gaping shells—unless they
chance to have closed upon any object in the water (e.g. another
Glochidium), in which case the valves remain firmly closed.”
In about four weeks after the Glochidium has quitted its host, and
the permanent shell has made its appearance within the two valves
of the Glochidium, the projecting teeth of the latter press upon the
ventral edge of the permanent shell, at a point about half way in its
lengthward measurement, retarding the growth of the shell at that
particular point, and indenting its otherwise uninterrupted curve with
an irregular notch or dent. As growth proceeds, this dent becomes
less and less perceptible on the ventral margin of the shell itself, but
its effects may be detected, in well-preserved specimens, by the
wavy turn in the lines of growth, especially near the umbones of the
young shell.
Mr. Latter found that all species of fish with which he
experimented had a strong dislike to Glochidium as an article of
food. Sometimes a fish would taste it “just to try,” but invariably spit it
out again in a very decided manner. The cause of unpleasantness
seemed not to be the irritation produced in the mouth of the fish by
the attempt of the Glochidium to attach itself, but was more probably
due to what the fish considered a nasty taste or odour in the object
of his attentions.

The following works will be found useful for further study of this
portion of the subject:—
F. M. Balfour, Comparative Embryology, vol. i. pp. 186–241.
F. Blochmann, Ueber die Entwickelung von Neritina fluviatilis Müll.:
Zeit. wiss. Zool. xxxvi. (1881), pp. 125–174.
L. Boutan, Recherches sur l’anatomie et le développement de la
Fissurelle: Arch. Zool. exp. gén. (2) iii. suppl. (1885), 173 pp.
W. K. Brooks, The development of the Squid (Loligo Pealii Les.):
Anniv. Mem. Bost. Soc. Nat. Hist. 1880.
„ „ The development of the oyster: Studies Biol. Lab.
Johns Hopk. Univ. i. (1880), 80 pp.
 R. von Erlanger, Zur Entwickelung von Paludina vivipara: Morph.
Jahrb. xvii. (1891), pp. 337–379, 636–680.
„ „ Zur Entwickelung von Bythinia tentaculata: Mitth. Zool.
Stat. Neap, x. (1892), pp. 376–406.
H. Fol, Sur le développement des Ptéropodes: Arch. Zool. exp.
gén. iv. (1875), pp. 1–214.
„ Etudes sur le développement des Mollusques.
Hétéropodes: ibid v. (1876), pp. 105–158.
„ Etudes sur le développement des Gastéropodes pulmonés:
ibid. viii. (1880), pp. 103–232.
H. Grenacher, Zur Entwickelungsgeschichte der Cephalopoden:
Zeit. wiss. Zool. xxiv. (1874), pp. 419–498.
B. Hatschek, Ueber Entwickelungsgeschichte von Teredo: Arb.
Zool. Inst. Univ. Wien, iii. (1881), pp. 1–44.
R. Horst, On the development of the European oyster: Quart.
Journ. Micr. Sc. xxii. (1882), pp. 339–346.
E. Korschelt and K. Heider, Lehrbuch der vergleichenden
Entwickelungsgeschichte der wirbellosen Thiere, Heft iii. (1893), pp.
909–1177 (the work is in process of translation into English).
A. Kowalewsky, Embryogénie du Chiton polii avec quelques
remarques sur le développement des autres Chitons: Ann. Mus. Hist.
Nat. Mars. Zool. i. (1883), v.
E. Ray Lankester, Contributions to the developmental history of
the Mollusca: Phil. Trans. Roy. Soc. vol. 165 (1875), pp. 1–31.
„ „ Observations on the development of the pond-snail
(Lymnaeus stagnalis), and on the early stages of other Mollusca:
Quart. Journ. Micr. Sc. xiv. (1874), pp. 365–391.
„ „ Observations on the development of the Cephalopoda:
ibid. xv. (1875), pp. 37–47.
W. Patten, The embryology of Patella: Arb. Zool. Inst. Univ. Wien,
vi. (1886), pp. 149–174.
M. Salensky, Études sur le développement du Vermet: Arch. Biol.
vi. (1885), pp. 655–759.
L. Vialleton, Recherches sur les premières phases du
développement de la Seiche (Sepia officinalis): Ann. Sc. Nat. Zool. (7)
vi. (1888), pp. 165–280.
 S. Watase, Observations on the development of Cephalopods:
Stud. Biol. Lab. Johns Hopk. Univ. iv. (1888), pp. 163–183.
„ „ Studies on Cephalopods: Journ. Morph. iv. (1891), pp.
247–294.
E. Ziegler, Die Entwickelung von Cyclas cornea Lam.: Zeit. wiss.
Zool. xli. (1885), pp. 525–569.
 CHAPTER VI
RESPIRATION AND CIRCULATION—THE MANTLE

The principle of respiration is the same in the Mollusca as in all

other animals. The blood is purified by being brought, in successive
instalments, into contact with pure air or pure water, the effect of
which is to expel the carbonic acid produced by animal combustion,
and to take up fresh supplies of oxygen. Whether the medium in
which a mollusc lives be water or air, the effect of the respiratory
action is practically the same.
Broadly speaking, Mollusca whose usual habitat is the water
‘breathe’ water, while those whose usual habitat is the land ‘breathe’
air. But this rule has its exceptions on both sides. The great majority
of the fresh-water Mollusca which are not provided with an
operculum (e.g. Limnaea, Physa, Planorbis), breathe air, in spite of
living in the water. They make periodic visits to the surface, and take
down a bubble of air, returning again for another when it is
exhausted. On the other hand many marine Mollusca which live
between tide-marks (e.g. Patella, Littorina, Purpura, many species of
Cerithium, Planaxis, and Nerita) are left out of the water, through the
bi-diurnal recess of the tide, for many hours together. Such species
invariably retain several drops of water in their branchiae, and, aided
by the moisture of the air, contrive to support life until the water
returns to them. Some species of Littorina (e.g. our own L. rudis and
many tropical species) live so near high-water mark that at neap-
tides it must frequently happen that they are untouched by the sea
for several weeks together, while they are frequently exposed to a
burning sun, which beats upon the rocks to which they cling. In this
case it appears that the respiratory organs will perform their
functions if they can manage to retain an extremely small amount of
moisture.[265]
The important part which the respiratory organs play in the
economy of the Mollusca may be judged from the fact that the
primary subdivision of the Cephalopoda into Dibranchiata and
Tetrabranchiata is based upon the number of branchiae they
possess. Further, the three great divisions of the Gasteropoda have
been named from the position or character of the breathing
apparatus, viz. Prosobranchiata, Opisthobranchiata and Pulmonata,
while the name Pelecypoda has hardly yet dispossessed
Lamellibranchiata, the more familiar name of the bivalves.
Respiration may be conducted by means of—(a) Branchiae or
Gills, (b) a Lung or Lung-cavity, (c) the outer skin.
In the Pelecypoda, Cephalopoda, Scaphopoda, and the great
majority of the Gasteropoda, respiration is by means of branchiae,
also known as ctenidia[266], when they represent the primitive
Molluscan gill and are not ‘secondary’ branchiae (pp. 156, 159).
In all non-operculate land and fresh-water Mollusca, in the
Auriculidae, and in one aberrant operculate (Amphibola), respiration
is conducted by means of a lung-cavity, or rarely by a true lung,
whence the name Pulmonata. The land operculates (Cyclophoridae,
Cyclostomatidae, Aciculidae, and Helicinidae) also breathe air, but
are not classified as Pulmonata, since other points in their
organisation relate them more closely to the marine
Prosobranchiata. Both methods of respiration are united in
Ampullaria, which breathes indifferently air through a long siphon
which it can elevate above the surface of the water, and water
through a branchia (see p. 158). Siphonaria (Fig. 57) is also
furnished with a lung-cavity as well as a branchia. Both these genera
may be regarded as in process of change from an aqueous to a
terrestrial life, and in Siphonaria the branchia is to a great extent
atrophied, since the animal is out of the water, on the average,
twenty-two hours out of the twenty-four. In the allied genus Gadinia,
where there is no trace of a branchia, but only a lung-cavity, and in
Cerithidea obtusa, which has a pulmonary organisation exactly
analogous to that of Cyclophorus,[267] this process may be regarded
as practically completed.
 Fig. 57.—A, Siphonaria gigas
Sowb., Panama, the
animal contracted in
spirit: gr, siphonal groove
on right side. B, Gadinia
peruviana, Sowb., Chili,
shell only: gr, mark of
siphonal groove to right
of head.
Respiration by means of the skin, without the development of any
special organ, is the simplest method of breathing which occurs in
the Mollusca. In certain cases, e.g. Elysia, Limapontia, and Cenia
among the Nudibranchs, and the parasitic Entoconcha and
Entocolax, none of which possess breathing organs of any kind, the
whole outer surface of the body appears to perform respiratory
functions. In others, the dorsal surface is covered with papillae of
varied size and number, which communicate with the heart by an
elaborate system of veins. This is the case with the greater number
of the Aeolididae (Fig. 58, compare Fig. 5, C), but it is curious that
when the animal is entirely deprived of these papillae, respiration
appears to be carried on without interruption through the skin.
 Fig. 58.—Aeolis despecta
Johnst., British coasts.
(After Alder and Hancock.)
In the development of a distinct breathing organ, it would seem as
if progress had been made along two definite lines, each resulting in
the exposure of a larger length of veins, i.e. of a larger amount of
blood, to the simultaneous operation of fresh air or fresh water.
Either (a) the skin itself may have developed, at more or less regular
intervals, elevations, or folds, which gradually took the form of
papillae, or else (b) an inward folding, or ‘invagination,’ of the skin, or
such a modification of the mantle-fold as is described below (p. 172)
may have taken place, resulting in the formation of a cavity more or
less surrounded by walls, within which the breathing organs were
ultimately developed. Sometimes a combination of both processes
seems to have occurred, and after a papilliform organ has been
produced, an extension or prolongation of the skin has taken place,
in order to afford a protection to it. Respiration by means of a lung-
cavity is certainly subsequent, in point of time, to respiration by
means of branchiae.

Fig. 59.—Chiton squamosus

L., Bermuda: A, anus;
Br, branchiae; M, mouth.

Fig. 60.—Fissurella virescens

Sowb., Panama, showing
position of the branchiae: Br,
branchiae: E, E, eyes; F, foot;
M, mantle; T, T, tentacles.
The branchiae seem to have been originally paired, and arranged
symmetrically on opposite sides of the body. It is not easy to decide
whether the multiple form of branchia which occurs in Chiton (Fig.
59), or the simple form as in Fissurella (Fig. 60), is the more
primitive. Some authorities hold that the multiple branchia has
gradually coalesced into the simple, others that the simple form has
grown, by serial repetition, into the multiple. There appears to be no
trace of any intermediate forms, and, as a matter of fact, the multiple
branchia is found only in the Amphineura, while one or rarely two
(never more) pairs of branchiae, occur, with various important
modifications, in the vast majority of the Mollusca.
Amphineura.—In Chiton the branchiae are external, forming a
long row of short plumes, placed symmetrically along each side of
the foot. The number of plumes, at the base of each of which lies an
osphradial patch, varies from about 70 to as few as 6 or 7. When the
plumes are few, they are confined to the posterior end, and thus
approximate to the form and position of the branchiae in the other
Amphineura. In Chaetoderma, the branchiae consist of two small
feather-shaped bodies, placed symmetrically on either side of the
anus, which opens into a sort of cloaca within which the branchiae
are situated. In Neomenia the branchiae are still further degraded,
consisting of a single bunch of filaments lying within the cloaca, while
in Proneomenia there is no more than a few irregular folds on the
cloaca-wall (Fig. 61).

Fig. 61.—Terminal portions of the Amphineura, illustrating the gradual degradation

of the branchiae, and their grouping round the anus in that class. A, Chiton
(Hemiarthrum) setulosus Carp., Torres Str.; B, Chiton (Leptochiton) benthus
Hadd., Torres Str.; C, Chaetoderma; D, Neomenia; a, anus; br, br, branchiae;
k, k, kidneys; p, pericardium. (A and B after Haddon, C and D after Hubrecht.)